Eur. J. Entomol. 107: 441–453, 2010 http://www.eje.cz/scripts/viewabstract.php?abstract=1553 ISSN 1210-5759 (print), 1802-8829 (online)

A new species group and five new species of the genus Pavania (: Dolichocybidae) associated with insects, with notes on leg chaetotaxy and the distribution of genera

HAMIDREZA HAJIQANBARl and ALEXNDER KHAUSTOV 2

1Department of Entomology, Faculty of Agriculture, Tarbiat Modares University, 14115-336, Tehran, Iran; e-mail: [email protected] 2 Nikita Botanical Gardens, National Scientific Center, Yalta, Crimea, 98648, Ukraine; e-mail: [email protected]

Key words. Acari, , Heterostigmata, Dolichocybidae, Pavania, taxonomy, new species, chaetotaxy, Coleoptera, Scarabaeidae, Carabidae, Iran

Abstract. A new species group and five new species of mites of the genus Pavania Lombardini, 1949 belonging to the family Doli- chocybidae (Acari: Heterostigmata) are described from Iran. Pavania gymnopleuri Hajiqanbar & Khaustov sp. n., P. sabzevarensis Hajiqanbar & Khaustov sp. n. and P. onthophagi Hajiqanbar & Khaustov sp. n., represent a new gymnopleuri species group. Two new species are also described in the fusiformis group: P. kamalii Hajiqanbar & Khaustov sp. n. and P. elongata Hajiqanbar & Khaustov sp. n. All new species are associated with scarabaeid and carabid beetles (Coleoptera: Scarabaeidae, Carabidae). This is the first record of the family Dolichocybidae from Iran. The leg setation and geographic distribution of all genera of this family are discussed and keys to genera and species of the genus Pavania are provided.

INTRODUCTION were collected directly in their habitats, i.e., livestock dung pads. In the section Heterostigmata, according to Mironov & The mites were cleared in lactophenol and mounted in Bochkov (2009), the basal superfamily Dolichocyboidea Hoyer’s medium. Morphology of the mites was studied using a includes the families Dolichocybidae Mahunka, 1970 and light microscope with phase contrast. All measurements in the Crotalomorphidae Lindquist & Krantz, 2002 (Lindquist descriptions are given in micrometers for the holotype and & Krantz, 2002). Despite this, these mites are morpho- (when available) for five paratypes (in parentheses). Termi- logically and biologically specialized. Mites of the family nology follows that of Lindquist (1986). Details of geographical Dolichocybidae are associated with different insects, coordinates were recorded using a GPS mobile device. The especially beetles and ants with which the female mites scarabaeid beetle hosts were identified by V. Martynov (Donetsk National University, Donetsk, Ukraine). have a phoretic relationship. Parasitic associations are not Holotypes of new taxa are deposited in the Acarological Col- recorded in this family. The adult females are fungivorous lection, Department of Entomology, Faculty of Agriculture, and demonstrate the phenomenon of neosomy (Kaliszew- Tarbiat Modares University, Tehran, Iran. One paratype of each ski et al., 1995) in which the larvae and males live within species is deposited in the following collections: Zoological the body of the physogastric females; males are therefore Institute, University of Hamburg, Hamburg, Germany; U.S. poorly known. National Museum of Natural History, Washington D.C., U.S.A.; Hitherto, this family consisted of two subfamilies, six Collection of Nikita Botanical Gardens, Yalta, Ukraine; Depart- genera and 30 species and was not recorded in Iran. ment of Animal Taxonomy and Ecology, A. Mickiewicz Uni- During an extensive survey of the insect-associated hete- versity, Poznan, Poland; and Acarological Collection, Zoo- rostigmatic mites in northeastern Iran, five new species of logical Museum, College of Agriculture, University of Tehran, Karaj, Iran. Most paratypes are retained with the holotype. the genus Pavania, belonging to the family Dolichocy- bidae, were recorded and are described here. In this SYSTEMATICS paper, we establish a new gymnopleuri species group Family Dolichocybidae Mahunka, 1970 containing 3 new species within the genus Pavania. Pavania is thus divided into two species groups: the fusi- Genus Pavania Lombardini, 1949 formis group (based on the name for the type species) Type species: Pavania fusiformis Lombardini, 1949 containing 15 species including two additional new spe- Diagnosis of adult female cies described below, and the gymnopleuri group con- Gnathosoma. Gnathosoma rounded subquadrangular, taining 3 species. usually longer than wide; dorsally with 2 pairs of cheli- MATERIAL AND METHODS ceral setae (ch1 and ch2) and ventrally with 1 pair of sub- capitular setae (su). Cheliceral stylets large and Mites were collected from different species of the beetle families Scarabaeidae and Carabidae (Insecta: Coleoptera). Of sickle-shaped. Pharynx hardly visible, rounded. Palpi the host beetles, some were captured in a light trap but most originated ventrolaterally from gnathosoma, 2-segmented.

441 Figs 1–2. Pavania gymnopleuri sp. n., female. 1 – body in dorsal view; 2 – body in ventral view. Scale bars 50 µm.

Palpal tibiotarsus with 2 solenidia and a small seta, termi- 4 (d, v’, v”, l’, M); Ta 6 (tc’, tc”, pv’, pv”, pl”, u’, Z). Leg nated in a distinct claw. Palpal femurogenu with 2 dorso- III: Fe 1 (d); Ge 1 (v’); Ti 4 (d, v’, v”, l’); Ta 5 (tc’, tc”, lateral setae (dGe and dFe). Small palpcoxal seta (pp) pv’, pv”, pl”). Leg IV: Fe 1 (d); Ge 1 (v’); Ti 4 (d, v’, v”, evident at palp bases. l’); Ta 5 (tc’, tc”, pv’, pv”, pl”). Idiosoma. Sometimes with tiny dimples on dorsal sur- Male unknown. face. Prodorsum with 3 pairs of setae (v1, v2 and sc2), The genus Pavania is here divided in two species setae v2 shorter than sc2 and v1; trichobothridia (sc1) pre- groups: sent or absent. Posterior edges of tergites undulated in Pavania gymnopleuri species group (Figs 1–18): Pro- some species. Cupuli ia and im obvious on tergites D and dorsum only with 3 pairs of setae (v1, v2 and sc2), tricho-

EF, respectively. Ventral apodemes usually well- bothridia (sc1) absent. developed; apodeme I (ap1) joined to the presternal apo- Pavania fusiformis species group (Figs 19–30): Pro- deme (appr) forming a Y-shaped structure. Each dorsum with 3 pairs of setae (v1, v2 and sc2) and one pair coxisternal plate with 3 pairs of setae; setae 3a and 3c of capitate trichobothridia (sc1). situated out of third coxisternal plate and inserted above Pavania gymnopleuri species group apodemes III. Genital plate with 2 pairs of setae (g1 and Pavania gymnopleuri Hajiqanbar & Khaustov sp. n. g2) and genital aperture. Aggenital plates spaced and flanking genital plate, and each bearing one seta (ag) and (Figs 1–6) a small porous area. Pseudanal plate with one pair of Differential diagnosis. The new species resembles two setae (ps) and anal slit. Legs. Leg I thicker than other other species of this species group: P. sabzevarensis and legs, legs II–IV similar in shape, 5-segmented, all with P. onthophagi. Readily recognizable from these two spe- empodium and paired claws, without spine-like setae and cies by the following characters: gnathosoma with cheli- all solenidia emergent. Number of setae and solenidia on ceral setae ch1 nearly 3 times longer than ch2 (1.7 times in each leg: Leg I: Fe 4 (d, v”, l’, l”); Ge 2 (v’, l’); Ti 6 (d, P. sabzevarensis and 1.5 times in P. onthophagi); dorsal k, v’, v”, l’, l”, M1, M2); Ta 11 (p’, p”, tc’, tc”, ft’, ft”, pl”, idiosomal setae smooth and pointed except c2 which is pv’, pv”, u’, u”, Z1, Z2). Leg II: Fe 2 (d, v”); Ge 1 (l’); Ti blunt-ended (c2, e and f blunt-ended in P. sabzevarensis

442 and most dorsal idiosomal setae weakly barbed and dis- tinctly blunt-ended in P. onthophagi); seta e longer than f (e shorter than f in P. sabzevarensis and e almost as long as f in P. onthophagi); seta h2 almost 15 times longer than h1 (7 times in P. sabzevarensis and nearly 3.5 times in P. onthophagi); ventral idiosomal setae smooth, attenuated and pointed, setae 2a and 2c longest on ventral aspect (ventral idiosomal setae smooth and needle-like, setae 2a longest on the ventral aspect of both P. sabzevarensis and P. onthophagi). Description Female (holotype). Length of idiosoma 144 (140–147 in 5 paratypes), width 86 (85–91). Gnathosoma (Figs 1–2). Length of gnathosoma 31 (28–31), width 28 (27–28). Gnathosomal capsule rounded subquadrangular in dorsal aspect, slightly longer than wide; cheliceral setae ch1 25 (21–23) and ch2 9 (9–12) pointed and smooth. Dorsomedian apodeme evident and forming the longitudinal division in dorsal view of gna- thosomal capsule. Cheliceral stylets 10 (9–11) long, curved and sickle-shaped. Subcapitulum with subcapi- tular setae su 7 (6–7) stiff, located on anterior half. Pharynx situated between subcapitular setae. Palpi con- spicuous, exceeding apex of gnathosomal capsule, femorogena with setae dGe 5 (5–7) and dFe 4 (4–5), tibi- otarsus with 2 solenidia, one longer than the other; minute palpcoxal setae pp inserted near to articulation of each palp with gnathosomal capsule. Idiosomal dorsum (Fig. 1). Ovoid in shape. All dorsal shields with smooth posterior margins, ornamented with sparse dimples, without backward undulating projections. Figs 3–6. Pavania gymnopleuri sp. n., female: legs I–IV in All dorsal idiosomal setae smooth. Prodorsal shield (PrS) dorsal view, respectively. Scale bar 20 µm. subtrapezoid, anterior margin approximately straight, pos- terior margin concave, with setae v1 35 (31–37), v2 5 subequal in size and slightly longer than 1c. Coxal field

(4–5), sc2 78 (77–84), setae v1 longer than distance II: 2a 27 (23–25), 2b 11 (9–10), 2c 30 (27–29), seta 2c between their bases, both v1 and sc2 tapered and pointed, longer than 2a and both more than 2 times longer than 2b. v2 very short and located posterolaterally to v1. Median Coxal field III: 3a 10 (8–9), 3b 10 (9–10), 3c 11 (10–11), shield of tergite C bearing setae c1 47 (45–46) tapered and all subequal in size. Coxal field IV: 4a 10 (9–11), 4b 12 pointed and each lateral shield with 1 distinctly blunt- (11–13), 4c 10 (10–12), setae 4a and 4c of the same ended seta c2 29 (28–33), c1 almost 1.6 times longer than length and shorter than 4b. Genital plate with setae g1 4 c2. Tergite D with setae d 50 (48–55) tapered and pointed (4–5), g2 5 (4–6). Aggenital plates each bearing seta ag and cupuli ia situated posterolaterally to setae d on each 10 (9–11) and 1 small porous area. Pseudanal plate with side, setae c1 and d subequal in length. Tergite EF with setae ps 5 (5–6). setae e 45 (38–43), f 27 (28–30) tapered and pointed, and Legs (Figs 3–6). Leg I (Fig. 3): setal formula (number 1 pair of cupuli im placed between insertion of setae e of solenidia in parentheses): 4-2-6(2)-11(2). Tarsus with and f, seta e nearly 1.7 times longer than f. Tergite H with 4 slightly blunt-ended eupathidial setae p’, p”, ft’ and ft”, setae h1 7 (6–8) needle-like and h2 108 (106–111) ultra- rest of setae on the leg I seta-like and pointed; pl” longest long, attenuated and pointed; seta h2 approximately 15 seta on tarsus; solenidion Z 6 (5-6) prominent, finger times longer than h1. Distances between dorsal idiosomal shaped and stalked with a fine apical spine; solenidion Z2 setae: v1–v1 30 (29–30), v2–v2 43 (40–41), v1–v2 6 (6–6), 3 (3-3) finger shaped. Tibia with eupathidium k stiff, sc2–sc2 49 (47–47), c1–c1 34 (33–36), d–d 62 (59–66), e–e thicker than other setae; solenidion M1 6 (6–7) fairly 46 (45–48), f–f 35 (36–38), e–f 16 (13–17), h1–h1 13 clubbed and stalked, longer than finger shaped M2 4 (3–4). (11–13), h2–h2 28 (28–29), h1–h2 7 (6–8). Genu with 2 setae (l’ and v’), l’ approximately twice as Idiosomal venter (Fig. 2). Ventral idiosomal setae long as v’. Seta d on femur longer than leg I; seta l” smooth, attenuated and pointed, setae 2a and 2c longest longer than v”. on ventral aspect. Apodemes II (ap2) not reaching to Leg II (Fig. 4). Setal formula: 2-1-4(1)-6(1). Setae of presternal apodeme (appr); both apodemes III (ap3) and leg attenuated and pointed except setae l’ on the genu and IV (ap4) not joined to poststernal apodeme (appo). Coxal v” on femur, which are needle-like, tc’ and tc” shortest field I: 1a 5 (5–5), 1b 5 (5–5), 1c 4 (4–5), 1a and 1b and longest setae on tarsus, respectively, solenidion Z 5

443 Figs 7–8. Pavania sabzevarensis sp. n., female. 7 – body in dorsal view; 8 – body in ventral view. Scale bars 50 µm.

(4–5) finger shaped, stalked and striated; solenidion M 4 Pavania sabzevarensis Hajiqanbar & Khaustov sp. n. (4–4) similar to Z but thinner; seta d on femur longest (Figs 7–12) seta of the leg II and 4 times as long as seta v”. Differential diagnosis. The new species differs from Leg III (Fig. 5). Setal formula: 1-1-4-5. Setae of leg two other species of this species group in its cheliceral attenuated and pointed except setae v’ on genu and d on setae ch1, which are nearly 1.7 times longer than ch2 (3 femur, which are needle-like; seta d of femur longer than times in P. gymnopleuri and 1.5 times in P. onthophagi); seta v’ of genu; setae d and v” subequal in length and seta dorsal idiosomal setae smooth and pointed except c2, e d longer than l’ on tibia. and f, which are blunt-ended (c2 blunt-ended in P. gymno- Leg IV (Fig. 6). Setal formula: 1-1-4-5. Setae of leg pleuri and most dorsal idiosomal setae weakly barbed and attenuated and pointed except setae v’ on genu and d on distinctly blunt-ended in P. onthophagi); seta e shorter femur, which are needle-like; seta v’ of genu longer than than f (e longer than f in P. gymnopleuri and e almost as seta d of femur; on tibia setae d and v” subequal in length long as f in P. onthophagi); seta h2 7 times longer than h1 and seta d more than twice as long as this segment. (almost 15 times in P. gymnopleuri and nearly 3.5 times Male unknown. in P. onthophagi); ventral idiosomal setae smooth and Type material. Holotype & (Sab-2-I) and 14& paratypes, needle-like, setae 2a the longest ones on the ventral phoretic on Gymnopleurus mopsus persianus Reitter, 1893 aspect (the same in P. onthophagi but ventral idiosomal (Coleoptera: Scarabaeidae) collected in dung pads of livestock: setae smooth, attenuated and pointed, setae 2a and 2c the Northeastern Iran, Razawi Khorasan province, Mehr village in longest on ventral aspect in P. gymnopleuri); genu I with the vicinity of Sabzevar town, 36.30°N, 57.80°E, altitude 1,024 one seta v’ (with two setae v’ and l’ in both P. gymno- m, 4.vi.2007, coll. H. Hajiqanbar. Etymology. The name gymnopleuri refers to the generic pleuri and P. onthophagi). name of the scarab host – Gymnopleurus.

444 Description Female (holotype). Length of idiosoma 126 (114–122 in 5 paratypes), width 83 (78–81). Gnathosoma (Figs 7–8). Length of gnathosoma 29 (29–30), width 25 (23–25). Gnathosomal capsule rounded subquadrangular in dorsal aspect, longer than wide; cheli- ceral setae ch1 15 (13–17) and ch2 9 (9–11) needle-like. Dorsomedian apodeme evident. Cheliceral stylets 9 (7–9) curved and falcate. Subcapitulum with subcapitular setae su 7 (6–7) stiff and located on anterior half. Palpi con- spicuous, exceeding apex of gnathosomal capsule, femorogenu with setae dGe 6 (6–7) and dFe 4 (4–5), tibi- otarsus with 2 solenidia, subequal in size; minute palp- coxal setae pp inserted near to articulation of each palp on gnathosomal capsule. Idiosomal dorsum (Fig. 7). Ovoid in shape. All dorsal shields ornamented with sparse dimples; some specimens with deep and large dimples limited to margins of pro- dorsum and tergites C and D; in some specimens poste- rior margins of some shields with backward undulating projections. All dorsal idiosomal setae smooth. Prodorsal shield subtrapezoid, with anterior margin curved and con- cave posterior margin, with setae v1 17 (15–17), v2 4

(4–5), sc2 45 (45–50), setae v1 shorter than distance between their bases, both v1 and sc2 tapered and pointed, v2 very short and located posterolaterally to v1. Median shield of tergite C including setae c1 32 (30–33) tapered and pointed and each lateral shield with 1 blunt-ended seta c2 25 (22–23), c1 approximately 1.3 times longer than c2. Tergite D bearing setae d 33 (30–34) tapered and pointed, and cupuli ia situated laterally to seta d, setae c1 Figs 9–12. Pavania sabzevarensis sp. n., female: legs I–IV in and d subequal in length, posterior margin of tergite D dorsal view, respectively. Scale bar 20 µm. with seven equally spaced tiny projections. Tergite EF with blunt-ended setae e 16 (15–17), f 18 (17–20) and 1 11 (10–11) and 1 small porous area. Pseudanal plate with pair of cupuli im placed anterolaterally to setae f, seta f setae ps 5 (5–6). slightly longer than e. Tergite H with setae h1 10 (10–11) Legs (Figs 9–12). Leg I (Fig. 9): setal formula: needle-like and h2 72 (72–77) ultra-long, attenuated and 4-1-6(2)-11(2). Tarsus with 4 slightly blunt-ended eupa- pointed; seta h2 almost 7 times as long as h1. Distances thidial setae p’, p”, ft’ and ft”, rest of setae on leg I between dorsal idiosomal setae: v1–v1 29 (26–29), v2–v2 slender and pointed; both tc” and pl” subequal in length 37 (35–38), v1–v2 4 (4–5), sc2–sc2 43 (40–43), c1–c1 30 and the longest on tarsus; solenidion Z1 5 (5–5) promi- (30–33), d–d 57 (57–58), e–e 44 (44–46), f–f 34 (33–36), nent, approximately spindle shaped; solenidion Z2 3 (3–4) e–f 10 (11–12), h1–h1 13 (11–12), h2–h2 26 (25–26), h1–h2 finger shaped. Tibia with eupathidium k stiff; solenidion

6 (5–7). M1 6 (7–8) stalked and slightly clubbed, longer than M2 4 Idiosomal venter (Fig. 8). All ventral idiosomal setae (3–4) finger shaped. Genu with 1 seta (v’). Seta d on smooth and needle-like; some specimens with deep and femur equal in length to whole leg I; seta v” longer than large dimples limited to margins of coxal fields. Apo- l”. demes II not reaching to presternal apodeme; both apo- Leg II (Fig. 10). Setal formula: 2-1-4(1)-6(1). Setae of demes III and IV not joined to the poststernal apodeme. leg attenuated and pointed except setae l’ on genu and v” Coxal field I with 1a 6 (6–7), 1b 5 (5–7), 1c 5 (5–6), both on femur, which are needle-like, pl” and tc” shortest and latter subequal in size and slightly shorter than 1a. Coxal longest setae on tarsus, respectively, solenidion Z 4 (3–4) field II with 2a 12 (10–12), 2b 7 (6–7), 2c 10 (9–11), seta finger shaped; solenidion M 3 (3–4) similar to Z but thin- 2a approximately 2 times as long as 2b. Coxal field III ner; seta d on femur 2.5 times as long as seta v”. with 3a 9 (9–10), 3b 9 (8–10), 3c 11 (10–11), setae 3a Leg III (Fig. 11). Setal formula: 1-1-4-5. Setae of leg and 3b sub-equal in length and both shorter than 3c. attenuated and pointed except setae pl” on the tarsus, v’ Coxal field IV with 4a 8 (8–9), 4b 11 (10–11), 4c 9 on genu and d on femur, which are needle-like; seta pl” (9–10), 4b longer than 4a and 4c. Genital plate without shortest on tarsus; seta d of femur as long as seta v’ of setae g1 and g2 (only their alveoli remain) but genital genu; seta v’ shorter than v” and d subequal to l’ on tibia. aperture visible. Aggenital plates each bearing 1 seta ag Leg IV (Fig. 12). Setal formula: 1-1-4-5. Setae of leg attenuated and pointed except setae pl” on the tarsus, v’

445 Figs 13–14. Pavania onthophagi sp. n., female. 13 – body in dorsal view; 14 – body in ventral view. Scale bars 50 µm. on genu and d on femur, which are needle-like; seta v’ of in P. gymnopleuri ventral idiosomal setae smooth, attenu- genu slightly longer than seta d on femur; on tibia setae d ated and pointed, setae 2a and 2c the longest in ventral shorter than v” and slightly longer than its segment. aspect). Male unknown. Description & & Type material. Holotype (Sab-2-II) and 21 paratypes Female (holotype). Length of idiosoma 136 (115–122 phoretic on Gymnopleurus mopsus persianus collected in dung in 5 paratypes), width 81 (74–77). pads of livestock: Northeastern Iran, Razawi Khorasan province, Mehr village in the vicinity of Sabzevar town, 36.30°N, Gnathosoma (Figs 13–14). Length of gnathosoma 28 57.80°E, altitude 1,024 m, 4.vi.2007, coll. H. Hajiqanbar. (28–30), width 28 (26–29). Gnathosomal capsule rounded Etymology. The specific epithet refers to the name of city subhexagonal in dorsal aspect, as long as wide with cheli- Sabzevar in Northeastern Iran. ceral setae ch1 18 (19–20) and ch2 12 (11–13) weakly barbed. Dorsomedian apodeme evident. Cheliceral stylets Pavania onthophagi Hajiqanbar & Khaustov sp. n. 8 (7–8) curved and falcate. Subcapitulum with subcapi- (Figs 13–18) tular setae su 5 (5–5) stiff, located on posterior half. Palpi Differential diagnosis. The new species is distinguish- somewhat appressed to gnathosoma (for this reason able from two other species of this species group by the accessories of tibiotarsus not visible), exceeding apex of following features: cheliceral setae ch1 approximately 1.5 gnathosomal capsule, femorogenu with setae dGe 9 (7–9) times longer than ch2 (3 times in P. gymnopleuri and 1.7 and dFe 5 (5–6) smooth and pointed; palpcoxal setae pp times in P. sabzevarensis); dorsal idiosomal setae weakly not visible. barbed and distinctly blunt-ended, except v2, sc2 and h2 Idiosomal dorsum (Fig. 13). Ovoid in shape. All being pointed (smooth in both other species, c2 blunt- dorsal shields ornamented with sparse and fairly deep ended in P. gymnopleuri and c2, e and f blunt-ended in P. dimples; in some specimens posterior margins of dorsal sabzevarensis); seta e almost as long as f (e longer than f shields with backward pointing and equally spaced pro- in P. gymnopleuri but shorter in P. sabzevarensis); seta h2 jections. Dorsal idiosomal setae weakly barbed and dis- nearly 3.5 times longer than h1 (almost 15 times in P. tinctly blunt-ended except v2, sc2 and h2 which are gymnopleuri and 7 times in P. sabzevarensis); ventral pointed. Prodorsal shield subtrapezoid, its anterior margin idiosomal setae smooth and needle-like, setae 2a the straight and posterior margin slightly concave, with setae longest in ventral aspect (the same in P. sabzevarensis but v1 17 (17–18), v2 7 (5–7), sc2 50 (47–52), setae v1 shorter

446 than distance between their bases, v2 short and located posterolaterally to v1, sc2 weakly barbed, tapered and pointed. Median shield of tergite C with setae c1 25

(23–25) and each lateral shield with seta c2 30 (28–30), c1 approximately 0.8 times longer than c2. Tergite D with setae d 27 (24–27) and cupuli ia situated posterior to seta d, seta d slightly longer than c1, posterior margin of ter- gite D usually undulated. Tergite EF with setae e 23 (21–23), f 23 (22–23) and 1 pair of cupuli im placed closer to setae e on its inner posterolateral side, seta f as long as e. Tergite H with setae h1 16 (15–17) and h2 57

(61–62), the latter long, attenuated and pointed, seta h2 almost 3.5 times longer than h1. Distances between dorsal idiosomal setae: v1–v1 28 (27–28), v2–v2 38 (35–36), v1–v2

4 (4–5), sc2–sc2 34 (33–35), c1–c1 25 (25–27), d–d 36 (36–39), e–e 35 (35–38), f–f 28 (27–29), e–f 8 (8–9), h1–h1 9 (10–10), h2–h2 22 (21–23), h1–h2 6 (5–6). Idiosomal venter (Fig. 14). All ventral idiosomal setae smooth and needle-like. Apodemes II almost reaching to presternal apodeme; both apodemes III and IV not joined with poststernal apodeme. Coxal field I: 1a 7 (6–7), 1b 5 (5–5), 1c 5 (5–5), 1c and 1b subequal in size and shorter than 1a. Coxal field II: 2a 12 (11–12), 2b 5 (5–6), 2c 11 (10–11), seta 2a 2.4 times as long as 2b. Coxal field III: 3a 9 (8–9), 3b 8 (8–9), 3c 8 (9–9), all setae in coxal field III almost subequal in length. Coxal field IV: setae 4a 7 (7–9), 4b 10 (9–10), 4c 9 (8–9), setae 4b longer than 4a and 4c. Genital plate with setae g1 3 (3–3) and g2 3 (3–3). Aggenital plates each bearing 1 seta ag 6 (6–7) and 1 small porous area. Pseudanal plate with setae ps 8 (8–9) and bifurcate anal slit. Legs (Figs 15–18). Leg I (Fig. 15): setal formula: 4-2-6(2)-11(2). Tarsus with 4 slightly blunt-ended eupa- Figs 15–18. Pavania onthophagi sp. n., female: legs I–IV in thidial setae p’, p”, ft’ and ft”, rest of setae on the leg I dorsal view, respectively. Scale bar 20 µm. seta-like and pointed; seta tc” longest on the tarsus and longer than pl”; solenidion Z1 6 (6–6) prominent and d of femur; on tibia setae d shorter than v” and longer finger shaped; solenidion Z2 3 (3–3) finger shaped. Tibia than its segment. Male unknown. with eupathidium k stiff; solenidion M1 7 (6–7) stalked and fairly clubbed, longer than M2 4 (4–5) finger shaped. Type material. Holotype & (Bo-1) and 15& paratypes pho- Genu with 2 setae (v’ and l’). Seta d on femur weakly retic on Onthophagus (Euonthophagus) gibbosus gibbosus barbed, shorter than leg I; seta l” longer than v”. (Scriba, 1970) (Coleoptera: Scarabaeidae) collected in dung Leg II (Fig. 16). Setal formula: 2-1-4(1)-6(1). Setae of pads of livestock: Northeastern Iran, North Khorasan province, leg attenuated and pointed except setae l’ on genu and v” Esfidan village in the vicinity of Bojnord town, 37.31°N, 57.57°E, altitude 1,611 m, 28.vi.2007, coll. H. Hajiqanbar. on femur which are needle-like; setae v” on tibia, l’ on Additional material. 11& (Es-1), phoretic on Onthophagus genu, d and v” on femur weakly barbed; seta tc’ shortest (s. str.) taurus (Schreber, 1959) (Coleoptera: Scarabaeidae) col- on tarsus, solenidion Z 4 (4–5) finger shaped; solenidion lected in dung pads of livestock: Northeastern Iran, North Kho- M 4 (4–5) similar to Z but thinner; seta d on femur 2.8 rasan province, vicinity of Esfarayen town, 36.93°N, 57.74°E, times as long as seta v”. altitude 1,471 m, 26.vi.2007, coll. H. Hajiqanbar. Leg III (Fig. 17). Setal formula: 1-1-4-5. Setae of leg Etymology. The specific epithet is derived from the generic attenuated and pointed except setae v’ on genu and d on name of the host scarab beetle – Onthophagus. femur which are needle-like; setae l’ on tibia, v’ on genu Pavania fusiformis species group and d on femur weakly barbed; seta pl” shortest on Pavania kamalii Hajiqanbar & Khaustov sp. n. tarsus; seta d of femur shorter than seta v’ of genu; setae d (Figs 19–24) shorter than v” and l’ on tibia. Leg IV (Fig. 18). Setal formula: 1-1-4-5. Setae of leg Differential diagnosis. This new species is similar to attenuated and pointed except v’ on genu and d on femur Pavania fusiformis Lombardini, 1949 but differs from it which are needle-like; setae l’ on tibia, v’ on genu and d by seta c1 longer than c2 (c2 longer than c1 in P. on femur weakly barbed; seta v’ of genu longer than seta fusiformis); seta c1 longer than d (c1 and d subequal in P.

447 Figs 19–20. Pavania kamalii sp. n., female. 19 – body in dorsal view; 20 – body in ventral view. Scale bars 50 µm.

fusiformis) and seta sc2 less than twice length of c1 (sc2 with setae v1 17 (17–19), v2 4 (4–5), sc2 39 (39–44) and 1 more than twice length of c1 in P. fusiformis). pair of barbed capitate trichobothridia sc1, setae v1 shorter Description than distance between their bases, v2 short and located lat- erally of v1, sc2 weakly barbed, tapered and pointed. Female (holotype). Length of idiosoma 130 (121–134 Median shield of tergite C with setae c1 28 (28–32) and in 5 paratypes), width 78 (71–75). each lateral shield with seta c2 22 (22–25), c1 almost 1.3 Gnathosoma (Figs 19–20). Length of gnathosoma 27 times longer than c2. Tergite D with setae d 21 (21–22) (26–29), width 25 (22–25). Gnathosomal capsule rounded and cupuli ia situated posterolaterally to seta d, seta c1 subquadrangular in dorsal aspect, longer than wide; dor- longer than d. Tergite EF with setae e 13 (13–15) and f 30 sally with cheliceral setae ch1 10 (11–14) and ch2 6 (7–8) (29–31), and 1 pair of cupuli im placed closer to setae f smooth. Dorsomedian apodeme evident. Cheliceral stylets on its inner anterolateral side, seta f 2.3 times as long as e. 9 (8–9) curved and falciform. Subcapitulum with subcapi- Tergite H with setae h1 12 (11–13) and h2 58 (61–65), the tular setae su 6 (6–7) needle-like. Pharynx rounded. Palpi latter long, attenuated and pointed, seta h2 almost 4.8 conspicuous, exceeding apex of gnathosomal capsule, times longer than h1. Distances between dorsal idiosomal femurogena with 2 setae dGe 7 (6–7) and dFe 4 (4–5), setae: v1–v1 22 (20–21), v2–v2 32 (30–32), v1–v2 5 (4–5), tibiotarsus with 2 solenidia subequal in size and 1 small sc2–sc2 34 (32–34), c1–c1 28 (25–28), d–d 52 (50–55), e–e seta; minute palpcoxal setae pp visible and inserted near 44 (43–45), f–f 33 (31–33), e–f 10 (10–11), h1–h1 11 to articulation of each palp with the gnathosomal capsule. (11–12), h2–h2 23 (23–25), h1–h2 6 (5–6). Idiosomal dorsum (Fig. 19). Almost ovoid in shape. Idiosomal venter (Fig. 20). All ventral idiosomal setae All dorsal shields smooth and their posterior margins with smooth and needle-like. Ventral apodemes well- several backward pointing and more or less equally developed; apodemes I joined with presternal apodeme; spaced projections. Dorsal idiosomal setae weakly barbed apodemes II not reaching to presternal apodeme; both and blunt–ended except v2 and h2, which are smooth and apodemes III and IV not joined to poststernal apodeme. pointed. Prodorsal shield subquadrangular, its anterior Coxal field I with setae 1a 7 (6–7), 1b 5 (5–6), 1c 5 margin curved and posterior margin slightly concave, (5–5), 1c and 1b subequal in length and shorter than 1a.

448 Coxal field II with setae 2a 11 (11–13), 2b 5 (5–6), 2c 10 (11–12), seta 2a 2.2 times longer than 2b. Coxal field III with setae 3a 8 (9–10), 3b 9 (9–10), 3c 9 (10–11), all setae of coxal field III subequal in length. Coxal field IV with setae 4a 9 (9–10), 4b 11 (11–12), 4c 10 (9–10), setae

4b longer than 4a and 4c. Genital plate with setae g1 2

(2–2), g2 3 (3–3) short. Aggenital plates including 1 seta ag 9 (9–11) and 1 small porous area each. Pseudanal plate with setae ps 6 (6–7). Legs (Fig. 21–24). Leg I almost thicker than leg II (Fig. 21): setal formula: 4-2-6(2)-11(2). Tarsus with 4 slightly blunt-ended eupathidial setae p’, p”, ft’ and ft”, rest of setae on leg I seta-like and pointed; seta tc” longest on tarsus and longer than pl”; solenidion Z1 6 (5–5) promi- nent and finger shaped; solenidion Z2 3 (3–3) finger shaped. Tibia with eupathidium k stiff; solenidion M1 7 (6–7) stalked and fairly clubbed, longer than M2 3 (3–4) finger shaped. seta d 4 times as long as l” on femur. Leg II (Fig. 22). Setal formula: 2-1-4(1)-6(1). Setae of leg attenuated and pointed; seta pl” shortest on tarsus, solenidion Z 4 (4–4) finger shaped; solenidion M 3 (3–3) similar to Z; on femur and genu, setae d and l’ subequal and both longer than seta v”. Leg III (Fig. 23). Setal formula: 1-1-4-5. Setae of leg attenuated and pointed; on the tarsus seta tc’ longer than its segment length; seta d of femur shorter than seta v’ of the genu. Leg IV (Fig. 24). Setal formula: 1-1-4-5. Setae of leg attenuated and pointed; on tarsus seta pl” shortest of all tarsals and shorter than its segment length; on tibia, seta l’ longer than d; seta v’ of genu longer than seta d on the femur. Figs 21–24. Pavania kamalii sp. n., female: legs I–IV in dorsal view, respectively. Scale bar 20 µm. Male unknown. & & Type material. Holotype (Es-3) and 11 paratypes pho- deme evident. Cheliceral stylets curved and falciform, 6 retic on Scarabaeus spp. (Coleoptera: Scarabaeidae) collected in long. Subcapitulum with subcapitular setae su 17 located dung pads of livestock or caught by a light trap: Northeastern on posterior half. Pharynx rounded. Palpi conspicuous, Iran, North Khorasan province, vicinity of Esfarayen town, 36.94°N, 57.73°E, altitude 1,483 m, 26. and 27.vi.2007, coll. H. exceeding apex of gnathosomal capsule, femurogenu with Hajiqanbar. setae dGe 11 and dFe 10, tibiotarsus with 2 solenidia, one Etymology. The new species is named in honour of Prof. longer than the other, and 1 small seta; minute palpcoxal Karim Kamali, supervisor of the senior author, in recognition of setae pp inserted near to articulation of each palp with his encouragement for the study of mites in Iran. gnathosomal capsule. Pavania elongata Hajiqanbar & Khaustov sp. n. Idiosomal dorsum (Fig. 25). Cylindrical in shape. All dorsal shields smooth and their posterior margins without (Figs 25–30) small projections. Dorsal idiosomal setae weakly barbed,

Differential diagnosis. The new species differs from attenuated and pointed except h2 which is smooth. Pro- other species of the species group by subequal coxal setae dorsal shield subquadrangular, its anterior margin slightly

2a and 2c (28) which are more than twice as long as 2b curved and posterior margin concave, with setae v1 28, v2

(13); seta c1 shorter than c2 but longer than d; seta f nearly 9, sc2 59 and 1 pair of smooth capitate trichobothridia sc1,

1.2 times as long as e. This species is also characterized setae v1 longer than distance between their bases, v2 short by having an elongate idiosoma (mites of the genus and located posterolaterally of v1. Median shield of tergite

Pavania usually have fusiform idiosoma). C with setae c1 26 and each lateral shield with seta c2 36 Description [one of the lateral shields is drawn from below due to rotation of the specimen during slide preparation], c2 Female (holotype). Length and width of idiosoma 200 almost 1.4 times longer than c1. Tergite D with setae d 23 and 85, respectively. and cupuli ia situated posterolaterally to seta d, seta c2 Gnathosoma (Fig. 25–26). Length of gnathosoma 28 longer than d. Tergite EF with setae e 20 and f 23, and 1 and 23, respectively. Gnathosomal capsule rounded pair of cupuli im placed close to setae f on its inner anter- subquadrangular in dorsal aspect, longer than wide; cheli- olateral side, seta f approximately 1.2 times as long as e. ceral setae ch1 21 and ch2 10 smooth. Dorsomedian apo-

449 Figs 25–26. Pavania elongata sp. n., female. 25 – body in dorsal view; 26 – body in ventral view. Scale bars 50 µm. Figs 27–30. Pavania elongata sp. n., female: legs I–IV in dorsal view, respectively. Scale bar 20 µm. Tergite H with setae h1 9 and h2 67 long, seta h2 almost

7.4 times longer than h1. Distances between dorsal idio- Seta d 2 times as long as l” on femur; both d and l” somal setae: v1–v1 14, v2–v2 21, v1–v2 4, sc2–sc2 28, c1–c1 weakly barbed. 40, d–d 55, e–e 52, f–f 38, e–f 23, h1–h1 11, h2–h2 27, Leg II (Fig. 28). Setal formula: 2-1-4(1)-6(1). Setae of h1–h2 7. Idiosomal venter (Fig. 26). All ventral idiosomal setae leg attenuated and pointed except d on femur which is smooth, tapered and pointed. Apodemes I joined to thin blunt-ended; setae l’ on tibia and genu and seta d on presternal apodeme; apodemes II reaching presternal apo- femur weakly barbed; setae pv” and pl” subequal in deme; both apodemes III and IV not joined to poststernal length and both the shortest setae on tarsus, solenidion Z apodeme. Coxal field I: 1a 14, 1b 16, 1c 12, 1b longer 4 finger shaped and fairly stalked; solenidion M 3 finger than1c and 1a. Coxal field II: 2a 28, 2b 13, 2c 28, setae shaped; on femur seta v” longer than d. 2a and 2c equal in length and the longest on ventral side. Leg III (Fig. 29). Setal formula: 1-1-4-5. Setae of leg Coxal field III: 3a 13, 3b 17, 3c 17, setae 3b and 3c equal attenuated and pointed; setae l’, v’ and v” on tibia, v’ on in length and longer than 3a. Coxal field IV: setae 4a 18, genu and d on femur weakly barbed; seta tc’ on tarsus 4b 18, 4c 16, setae 4a and 4b subequal in length and shorter than its segment length; seta d of femur almost as longer than 4c. Genital plate with subequal short setae g1 long as seta v’ of genu.

5, g2 5. Aggenital plates including 1 seta ag 19 and 1 Leg IV (Fig. 30). Setal formula: 1-1-4-5. Setae on leg small porous area each. Pseudanal plate with setae ps 9. attenuated and pointed; setae l’ and v” on tibia, v’ on the Legs (Figs 27–30). Leg I thicker and longer than other genu and d on femur weakly barbed; seta pl” on tarsus legs (Fig. 27): setal formula: 4-2-6(2)-11(2). Tarsus with longer than its segment length; seta l’ on tibia approxi- 4 slightly blunt-ended eupathidial setae p’, p”, ft’ and ft”, mately as long as d; seta v’ of genu and seta d of femur rest of setae on leg I slender and pointed; seta pl” longest subequal in length. Male unknown. on tarsus and longer than tc”; solenidion Z1 5 prominent, slightly stalked and finger shaped; solenidion Z2 4, finger Type material. Holotype & (Ms-6) phoretic on an unidenti- shaped. Tibia with eupathidium k; solenidion M1 6 stalked fied carabid beetle (Coleoptera: Carabidae) collected on surface of the ground: Northeastern Iran, North Khorasan province, and distinctly clubbed, longer than M2 4 finger shaped. Haver village, elevations of Maneh and Samalghan town,

450 Fig. 31. Distribution of the dolichocybid genera in the world. y – Pavania; M – Dolichocybe; ­ – Rafamastix; 3 – Acanthomastix; 9 – Dolichomotes; ~ – Formicomotes.

37.7°N, 56.4°E, altitude 2,365 m, 30.vi.2006, coll. H. Hajiqan- Features of the recently discovered taxa (after 1980) bar. suggest that the previously defined characters of these Etymology. The specific epithet is derived from the Latin subfamilies are not convenient and constant. On the one root of “elongatus” meaning prolonged, which refers to the hand, Rafamastix contravenes the number of setae on elongate idiosoma of this species. coxal fields III and IV in the subfamily Formicomotinae, DISCUSSION and on the other, the new Pavania gymnopleuri species group invalidates the presence of trichobothridia as a con- Sevastianov (1980) classified five genera of the family stant character of the subfamily Dolichocybinae. There- Dolichocybidae in the two subfamilies Formicomotinae fore, additional characters are needed to define the and Dolichocybinae, based on the presence or absence of subfamilies. Leg setation of all genera of the family Doli- trichobothridia and numbers of setae on the coxal fields. chocybidae were studied (see Table 1). This revealed He considered absence of trichobothridia and presence of some useful characters that were ignored in the past one pair of setae on each coxal field as features defining owing to the often imperfect description of leg setation at the subfamily Formicomotinae, which included the gene- the generic level. ra Dolichomotes Smiley, 1967, Acanthomastix Mahunka, It should be noted that the solenidia in genera Dolicho- 1970 and Formicomotes Sevastianov, 1980. He placed motes and Formicomotes are attached in a somewhat the genera Pavania Lombardini, 1947 and Dolichocybe unusual way. All solenidia in these genera are inserted Krantz, 1957 in the subfamily Dolichocybinae, character- into the surface of the leg segments, except 1 on first ized by the presence of the trichobothridia and more than M tibia, which has a normal structure, emergent and stalked. one and up to four pairs of setae on the coxal fields. In this regard, Dolichomotes crossi is an exception in that Later on, Magowski (1997) described the genus Rafa- mastix and placed it in the subfamily Formicomotinae. In solenidion Z2 is not pustulate but rather emergent and mites of this genus, however, coxal fields III and IV each stick-like. bear two pairs of setae. According to the summarized data in Table 1, some The gymnopleuri species group of genus Pavania additional features (putative apomorphies), originating described above is morphologically and ecologically very from leg setation, can be proposed for improving the similar to the fusiformis group. Even the leg setation in diagnoses of the two subfamilies. these two groups is identical (Figs 3–6 and 21–24). Like- Dolichocybinae: solenidion M on tibia III absent (i), wise, we recorded species of the gymnopleuri group pho- solenidion M on tibia IV absent (ii). retic on scarab beetles, which are also recorded as hosts Formicomotinae: setae v” and l’ on femur I absent (i), of the fusiformis species group. The new species group seta v’ on genu I absent (ii), setae u’ and u” on tarsus I differs from the fusiformis group by a single probably absent (iii), seta u’ on tarsus II absent (iv), seta tc’ on apomorphic feature – loss of trichobothridia. The fusi- tarsus IV absent (v). formis group is not defined by a distinct apomorphy and The family now includes six genera and 35 species. The may be a paraphyletic stem group. Thus, the genus genus Pavania, including the five new species described, Pavania belongs obviously to the subfamily Dolichocy- includes 18 species and constitutes the largest genus of binae regardless of the lack of trichobothridia (being one the family. The remaining 17 species are distributed as of Sevastianov’s criteria for diagnosis of the subfamily follows: Dolichocybe (8 species), Acanthomastix (4), Formicomotinae) in the newly defined gymnopleuri Dolichomotes (2), Formicomotes (2) and Rafamastix (1). group.

451 TABLE 1. Leg setation of the family Dolichocybidae (females). The newly described taxa expanded the geographical * distribution of the dolichocybid mites in the Palaearctic Region. Fig. 31 shows map of the distribution of the genera of the family. All six zoogeographic realms har- bour at least one dolichocybid genus. There is only one Pavania Dolichocybe Rafamastix Acanthomastix Dolichomotes Formicomotes Leg I genus in each of the Afrotropical and Australasian realms, Femur d d d – – – Pavania and Acanthomastix, respectively. There are two l” l” l” l” l” l” genera in the Neotropical realm, Pavania and Rafamastix, v” v” – – – – the latter being restricted to that realm. There are three l’ l’ – – – – Genu v’ v’ – – – – genera in both the Oriental and Nearctic realms; Dolicho- l’** l’ l’ l’ l’ l’ cybe, Dolichomotes and Formicomotes in the Oriental, Tibia M1 M1 M1 M1 M1 M1 Formicomotes is replaced by Acanthomastix in the Nearc- M2 M2 M2 M2 M2 M2 k k k k k k tic. Finally, four genera and most of the dolichocybid d d d d d d mites occur in the Palaearctic realm. l” l” l” l” l” l” v” v” v” v” v” v” Key to world genera of the family Dolichocybidae v’ v’ v’ – – – (females) l’ l’ l’ l’ l’ l’ 1 Each coxal field with 3 pairs of setae; solenidion on tibiae Tarsus Z1 Z1 Z1 Z1 Z1 Z1 Z2 Z2 Z2 Z2 Z2 Z2 III–IV absent (subfamily Dolichocybinae)...... 2 p’ p’ – p’ p’ p’ – Each coxal field with less than 3 pairs of setae; solenidion p” p” – – – p” on tibiae III–IV present (subfamily Formicomotinae)..... 3 tc’ tc’ tc’ tc’ tc’ tc’ tc” tc” tc” tc” tc” tc” 2 Gnathosoma longer than wide; small and indis- ft’ ft’ ft’ ft’ – – tinct; tarsus I with 10 setae (pl” absent); with deep constric- ft” ft” ft” ft” – – tion between propodosoma and hysterosoma which are pv’ pv’ pv’ pv’ pv’ pv’ separated by soft and transversely striated cuticle...... pv” pv” pv” pv” pv” pv” pl” – – pl” pl” pl” ...... Dolichocybe u’ u’ – – – – – Gnathosoma hardly as long as wide; chelicerae large and u” u” – – – – distinct; tarsus I with 11 setae (pl” present); without deep Leg II constriction between propodosoma and hysterosoma...... Femur d d d – – – ...... Pavania v” v” l” l” l” l” 3 Coxal fields III–IV with 2 pairs of setae each. . . Rafamastix Genu l’ l’ l’ l’ l’ l’ – Coxal fields III–IV with 1 pair of setae each...... 4 Tibia M M M M M M d d d d d d 4 All solenidia emergent; femora III–IV each with one seta v” v” v” v” v” v” (l”); tarsus I with both fastigial setae (ft’, ft”)...... v’ v’ v’ – – – ...... Acanthomastix l’ l’ l’ l’ l’ l’ – Most solenidia joined to surface of leg and pustulate; femora Tarsus Z Z Z Z Z Z tc’ tc’ tc’ tc’ tc’ tc’ III–IV without seta; tarsus I without fastigial setae...... 5 tc” tc” tc” tc” tc” tc” 5 Tarsus III with 4 setae (tc’ absent); tarsus I with 6 setae (p” pv’ pv’ pv’ pv’ pv’ pv’ absent)...... Dolichomotes pv” pv” pv” pv” pv” pv” – Tarsus III with 5 setae (tc’ present); tarsus I with 7 setae (p” pl” pl” pl” pl” pl” pl” u’ u’ – – – – present)...... Formicomotes Leg III Key to world species of the genus Pavania (females) Femur d d l’ (?) l” – – 1 Trichobothridia absent (gymnopleuri species group)...... 2 Genu v’ v’ l’ l’ l’ l’ – Trichobothridia present (fusiformis species group)...... 4 Tibia – – M M M M d d d d d d 2 Setae v1 longer than distance between their bases; setae ch1 3 v” v” v” v” v” v” times longer than ch2; setae e longer than f; setae h2 15 times v’ v’ v’ – – – longer than h1 (Iran)...... l’ l’ l’ l’ l’ l’ ...... P. gymnopleuri Hajiqanbar & Khaustov sp. n. Tarsus tc’ tc’ tc’ tc’ – tc’ tc” tc” tc” tc” tc” tc” – Setae v1 shorter than distance between their bases; setae ch1 pv’ pv’ pv’ pv’ pv’ pv’ less than 3 times longer than ch2; setae e never longer than f; pv” pv” pv” pv” pv” pv” setae h2 at most 7 times longer than h1...... 3 pl” pl” pl” pl” pl” pl” 3 Genu I with one seta (v’); dorsal idiosomal setae smooth; Leg IV setae c1 longer than c2; setae c1 and d pointed (Iran)...... Femur d d l’ (?) l” – – ...... P. sabzevarensis Hajiqanbar & Khaustov sp. n. Genu v’ v’ l’ l’ l’ l’ – Genu I with two setae (v’, l’); dorsal idiosomal setae weakly Tibia – – M M M M d d d d d d barbed; setae c2 longer than c1; setae c1 and d distinctly v” v” v” v” v” v” blunt-ended (Iran)...... v’ v’ v’ – – – ...... P. onthophagi Hajiqanbar & Khaustov sp. n.

l’ l’ l’ l’ l’ l’ 4 Setae h2 less than 3.5 times longer than h1...... 5 Tarsus tc’ tc’ – – – – – Setae h more than 3.5 times longer than h ...... 12 tc” tc” tc” tc” tc” tc” 2 1 pv’ pv’ pv’ pv’ pv’ pv’ 5 Setae c1 reaching beyond bases of setae f; setae c1 longer pv” pv” pv” pv” pv” pv” than h2; setae d and h2 subequal (Ghana)...... pl” pl” pl” pl” pl” pl” ...... P. perhirsuta Mahunka, 1973 * based on D. crossi; ** lacking in P. sabzevarensis sp. n.

452 – Setae c1 never reaching beyond bases of setae f; setae c1 16 Setae e and f subequal; setae e longer than h1; setae on epi-

shorter than h2; setae d shorter than h2...... 6 meres II subequal (Tajikistan)......

6 Setae h1 longer than or subequal to d, e and f ...... 7 ...... P. tadjikistanica Sevastianov, 1980

– Setae h1 shorter than d, e and f ...... 11 – Setae f more than twice as long as e; setae e and h1 subequal;

7 Setae h1 longer than sc2; setae sc2 distinctly shorter than dis- setae on epimeres II not subequal...... 17 tance between their bases; seta e inserted at the same level 17 Most dorsal idiosomal setae weakly barbed and blunt-ended;

as seta f (Ghana)...... P. equisetosa Mahunka, 1975 setae c1 longer than c2; setae sc2 less than twice as long as c1

– Setae h1 shorter than sc2; setae sc2 longer than or subequal to (Iran)...... P. kamalii Hajiqanbar & Khaustov sp. n. distance between their bases; seta f inserted posteriorly to – Dorsal idiosomal setae smooth and pointed; setae c2 longer seta e...... 8 than c1; setae sc2 more than twice as long as c1 (Italy)......

8 Setae sc2 distinctly longer than distance between their bases; ...... P. fusiformis Lombardini, 1949

setae sc2 reaching beyond bases of setae c1...... 9 ACKNOWLEDGEMENTS. We are indebted to W.L. – Setae sc2 subequal to distance between their bases; setae sc2 Magowski (Department of Animal Taxonomy and Ecology, A. never reaching beyond bases of setae c1...... 10 Mickiewicz University, Poznan, Poland) for reviewing the 9 Setae c1 longer than c2; setae h1 longer than e; setae h1 and v1 subequal (Ukraine, Slovakia). . P. riparia Sevastianov, 1980 manuscript and his valuable and helpful comments. We also express our gratitude to V. Martynov (Donetsk National Univer- – Setae c2 longer than c1; setae h1 and e subequal; setae h1 sity, Donetsk, Ukraine) for identifying the scarabaeid beetles. longer than v1 (Ghana)...... P. luisiae Mahunka, 1974 Special thanks go to A. Khurani (Department of Geography, 10 Setae c1, d, e and f blunt-ended; setae v1 reaching beyond Faculty of Humanity, Tarbiat Modares University, Tehran, Iran) bases of setae ch1 (Russia)...... P. carabidophila Khaustov, 2005 for providing the distribution map of the genera, which was obtained using GIS software. – Setae c1, d, e and f pointed; setae v1 never reaching beyond bases of setae ch (Ukraine)....P. bembidii Khaustov, 2005 1 REFERENCES 11 Setae h2 more than twice as long as h1; setae e and f sube-

qual and both longer than d; setae f distinctly longer than c1 KALISZEWSKI M., ATHIAS-BINCHE F. & LINDQUIST E.E. 1995: Para- (Egypt)..... P. tahanae Sevastianov and Abo-Korah, 1985 sitism and parasitoidism in Tarsonemina (Acari: Heterostig-

– Setae h2 less than twice as long as h1; setae e and d subequal mata) and evolutionary considerations. Adv. Parasitol. 35: and both longer than f; setae c1 and f subequal (Russia, 335–367. Turkmenistan, Iran)...... P. protracta Sevastianov, 1980 LINDQUIST E.E. 1986: The world genera of Tarsonemidae (Acari:

12 Setae h2 more than 6 times longer than h1...... 13 Heterostigmata): A morphological, phylogenetic and system-

– Setae h2 less than 6 times longer than h1...... 16 atic revision, with a reclassification of family-group taxa in

13 Setae sc2 at least 3.5 times longer than v1; setae f more than Heterostigmata. Mem. Entomol. Soc. Can. 136: 1–517. twice as long as e; setae e longer than v1 (Ghana)...... LINDQUIST E.E. & KRANTZ G.W. 2002: Description of, and vali- ...... P. endroedyi Mahunka, 1975 dation of names for the genus Crotalomorpha and the family

– Setae sc2 less than 2.5 times longer than v1; setae f less than Crotalomorphidae (Acari: Heterostigmata). Syst. Appl. twice as long as e; setae e shorter than v1...... 14 Acarol. 7: 129–142. 14 Setae sc2 less than twice as long as v1; setae f longer than d; MAGOWSKI W. 1997: Rafamastix plenus gen. n., sp. n. of the setae c1 reaching beyond posterior border of tergite C (Bra- family Dolichocybidae from South America (Acari: Hetero- zil)...... P. brasiliensis Mahunka, 1970 stigmata). Genus 8: 61–74. – Setae sc2 more than twice as long as v1; setae f and d sube- MIRONOV S.V. & BOCHKOV A.V. 2009: Modern conceptions con- qual; setae c1 never reaching beyond posterior border of ter- cerning the macrophylogeny of acariform mites (, gite C...... 15 ). Zool. Zh. 88: 922–937 [in Russian]. 15 Setae 2a as long as 2c and both longer than c1, d and f; setae SEVASTIANOV V.D. 1980: New taxa of mites of the family Doli- su protruding beyond anterior border of gnathosoma (Iran). . chocybidae (Trombidiformes, Tarsonemina) and phylogenetic ...... P. elongata Hajiqanbar & Khaustov sp. n. relations of its subfamilies. Zool. Zh. 59: 1453–1462 [in Rus- – Setae 2a longer than 2c and both shorter than c1, d and f; sian]. setae su never protruding beyond anterior border of gnatho- soma (Ghana)...... P. simplex Mahunka, 1973 Received August 20, 2009; revised and accepted February 17, 2010

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